Lymantria monacha (nun moth)
- Summary of Invasiveness
- Taxonomic Tree
- Notes on Taxonomy and Nomenclature
- Distribution Table
- History of Introduction and Spread
- Habitat List
- Hosts/Species Affected
- Host Plants and Other Plants Affected
- Growth Stages
- List of Symptoms/Signs
- Biology and Ecology
- Natural enemies
- Notes on Natural Enemies
- Means of Movement and Dispersal
- Plant Trade
- Impact Summary
- Environmental Impact
- Social Impact
- Detection and Inspection
- Similarities to Other Species/Conditions
- Prevention and Control
- Distribution Maps
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PicturesTop of page
IdentityTop of page
Preferred Scientific Name
- Lymantria monacha (Linnaeus)
Preferred Common Name
- nun moth
Other Scientific Names
- Bombyx eremita Hübner, 1808
- Bombyx nigra Freyer, 1833
- Liparis monacha Linnaeus
- Liparis monacha var. oethiops De Selys-Longchamps, 1857
- Lymantria brunnea Stipan, 1933
- Lymantria fasciata Hannemann, 1916
- Lymantria kusnezovi Kulossow, 1928
- Lymantria monacha chosenibia Bryk
- Lymantria monacha eremita
- Lymantria monacha flaviventer Kruilikovsky
- Lymantria monacha gracilis Kruilikovsky
- Lymantria monacha idae Bryk
- Lymantria monacha lateralis Bryk
- Lymantria monacha matuta Bryk
- Lymantria monacha nigra
- Lymantria transiens Lambillion, 1909
- Noctua heteroclita Müller, 1764
- Ocneria monacha Linnaeus
- Phalaena Bombyx monacha Linnaeus, 1758
- Phalaena monacha Linnaeus
- Porthetria monacha Linnaeus
- Psilura monacha Linnaeus
- Psilura transiens Thierry Mieg, 1886
International Common Names
- English: black arched tussock moth; black arches moth; tussock moth
- Spanish: lagarta monacha; mariposa monacha; mariposa monja; monja
- French: bombix moine; bombyx moine; moine; nonne
- Russian: monashenka; shelkopryad monashenka
Local Common Names
- Austria: nonne
- Czech Republic: bekyne mniska
- Denmark: nonnen
- Finland: havununna
- Germany: fichten spinner; nonne
- Italy: monaca
- Japan: nonne-maimai
- Korea, DPR: eolrukmaemi-nabang
- Netherlands: nonnetje; nonvlinder
- Norway: barskognonne
- Poland: brudnica mniszka
- Sweden: barrskogsnunna
- LYMAMO (Lymantria monacha)
Summary of InvasivenessTop of page
L. monacha is considered to be the number one forest pest in Poland because of the unprecedented economic losses it causes in spite of intensive chemical protective treatments on an area of 6.3 million ha of pine, spruce and other conifers between 1978 and 1984 (Sliwa and Sierpinski, 1986). It is also considered to be a major pest in all the other areas where it goes through periodic outbreaks causing defoliation and resulting in the death of spruce and pine trees (Bejer, 1988). The frequency of outbreaks has increase from about every 30 to 40 years to intervals of 6 years. It poses an ever present threat of being accidentally transported via commerce and introduced into other world areas, where susceptible hosts are present. This is because the adults are readily attracted to artificial lights and have been observed in Russian Far East ports (Munson et al., 1995), and although the eggs are normally laid in bark crevices, they could also be deposited in crevices on containers, pallets, ships, etc. In a pest risk assessment for importation of larch from Siberia into the USA, L. monacha was one of the serious pests that were considered at risk of introduction if the bark was still on the logs, because of their use of the bark for oviposition and the fact that the eggs are not readily visible (Anonymous, 1991). Its establishment in areas with suitable hosts would be disastrous because of its polyphagous feeding habits, ability to colonize new habitats, and capacity to be spread rapidly by flying females.
L. monacha is listed as an invasive species of concern by the United States Department of Agriculture (USDA); port inspectors monitor for it and as part of the Rapid Detection Pilot Project pheromone traps are being placed near ports of entry to detect any breeding populations. There were no L. monacha trapped in the one season of USA port monitoring reported on so far [at the time of writing in 2004]. Population levels are being monitored through collaboration between the USA and Russian agencies in the Russian Far East near ports (Munson et al., 1995).
Taxonomic TreeTop of page
- Domain: Eukaryota
- Kingdom: Metazoa
- Phylum: Arthropoda
- Subphylum: Uniramia
- Class: Insecta
- Order: Lepidoptera
- Family: Erebidae
- Subfamily: Lymantriinae
- Genus: Lymantria
- Species: Lymantria monacha
Notes on Taxonomy and NomenclatureTop of page
DescriptionTop of page
The eggs of L. monacha are spherical, approximately 1 mm in diameter, slightly depressed in the middle of the upper surface, and often flattened on the lower surface. The eggs are orange-brown (blue-green if reared on an artificial diet) at first, and later turn brown with an opalescent shine. The eggs are deposited in clumps that are glued together without a covering of hair. The female does not deposit all of her 70 to 300 eggs in one place and generally hides them in crevices in the bark of trees.
The hairy larvae of the Lymantriidae can always be distinguished from other families by the presence of some type of dorsal eversible glands, prominently located in the middle of the sixth and seventh abdominal segments. The larvae of Lymantria species have a full complement of low, rounded verrucae, without dense hair tufts, and usually without hair pencils. The dorsal verrucae bear needle-like setae and sometimes longer hairs.
Newly hatched larvae are approximately 4 mm long. At first, they appear tan-coloured but within several hours they turn black. They are very hairy and have 'air hairs', which may aid in dispersal. The 'air hairs' are simple setae with a bulb-like structure in the middle that looks like a water droplet under the light microscope. The 'air hairs' are only present on the first-instar larvae.
The second-instars appear black with a few lighter spots and have two white patches that almost encircle each dorsal verrucae on the third thoracic segment, but do not meet along the mid-dorsal line. There is also a light patch that fills the mid-dorsal space between the verrucae from the middle of the fourth to the middle of the sixth abdominal segments. The larvae have small, paired glands on the first and fifth abdominal segments and large, single orange eversible glands on the sixth and seventh, which are clearly visible.
From the third-instar on, the head of the larva is orange-brown with numerous brown and black freckles. The mid-dorsal stripe is a mottled brown to black. The dorsal verrucae of the larva are all bluish. The dorsal spot or patch patterns present in the second-instar persist through the later instars. The mature larvae appear tanish, greenish or dark-greyish, with extensive brown or black mottling and are 30 to 40 mm long. The colour of the larvae conceals them when they rest on the branch of a conifer.
The pupa has no cocoon, is reddish-brown, and shiny with light-coloured (occasionally red) clumps of hairs. It is 18 to 25 mm long. The sex of the pupa can be determined by the form of the bases of the antennal pads and by the characteristics of the future sex organs, located on the external ventral portion of the abdominal segments (female on the eighth segment and male on the ninth).
Lymantrid adults can usually be recognised by the position of the Sc vein relative to the Rs vein in the hindwing; the base of the M2 vein being much closer to M3 than to M1 in the hindwing; the absence or vestigial nature of the haustellum; the absence of ocelli; the prespiracular counter tympanal hood; and the one to three long, divergent spinules at the end of each antennal branch (Ferguson, 1978).
In Lymantria species, the females have wings that are longer and narrower than those of the male. The female bodies are stout and the antennae are bipectinate, with short branches approximately the thickness of the shaft, each bearing one terminal spinule. The male antennae are bipectinate with very long branches, each bearing one long terminal spinule and sometimes a second very short one. Sexual dimorphism in form and colour is often extreme. When at rest, the outline of the female resembles an isosceles triangle, whereas that of the male resembles an equilateral triangle.
Sexual dimorphism in colour is much reduced in L. monacha. The forewing coloration of both sexes varies from characteristic chalk-white, decorated with numerous dark transverse wavy lines and patches, to almost black. The hind wings are generally grey-brown with minute dark and/or light patches at their edge. The female has a wingspan of 45 to 55 mm, whereas the male has a wingspan of 35 to 45 mm. Lighter coloured females have abdomens with patches of pink or red and black bands, which correspond to the intersegments. Darker coloured females have abdomens that are all dark. The darker forms are common in Europe but totally absent in Oriental populations. The female has an extremely long ovipositor adapted for its specialised egg-laying habit.
DistributionTop of page
L. monacha is of Eurasian origin. See the Biology and Ecology/Environmental Requirements section for the limits of distribution. The most frequent outbreaks appear to occur in the Poland/Germany region of Europe (Lipa and Glowacka, 1995). The earliest reported outbreaks occurred in Poland in 1783-1784, 1794-1798, 1892-1898 and Western Russia (presently Estonia, Latvia, and Lithuania) in 1827-1829 (Wellenstein, 1942a; Sliwa, 1987). The history of outbreaks throughout Eurasia is summarized by Wellenstein (1942a), Marushina (1978), Bejer (1988), Schönherr (1989), and Lipa and Glowacka (1995).
Distribution TableTop of page
The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.Last updated: 30 Jun 2021
|Continent/Country/Region||Distribution||Last Reported||Origin||First Reported||Invasive||Reference||Notes|
|-Guizhou||Present||Native||Invasive||Original citation: Chao (1978)|
|-Sichuan||Present||Native||Invasive||Original citation: Sun (1989)|
|-Xinjiang||Present||Native||Invasive||Original citation: Chao (1978)|
|-Kyushu||Present||Native||Invasive||Original citation: Inoue (1957)|
|-Ryukyu Islands||Absent, Formerly present||Original citation: Kishida (1987)|
|-Shikoku||Present||Native||Invasive||Original citation: Inoue (1957)|
|Kazakhstan||Present||Native||Invasive||Original citation: Gninenko, 1993|
|Taiwan||Absent, Formerly present||Original citation: Kishida (1987)|
|Bosnia and Herzegovina||Present|
|Estonia||Present||Native||Invasive||Original citation: Marushina (1978)|
|Latvia||Present||Native||Invasive||Original citation: Marushina (1978)|
|Lithuania||Present||Native||Invasive||Original citation: Marushina (1978)|
|North Macedonia||Present||Native||Invasive||Original citation: Dzutevski and Cakar (1955)|
|-Central Russia||Present, Widespread||Native||Invasive||Original citation: Marushina (1978)|
|-Eastern Siberia||Present, Widespread||Native||Invasive||Original citation: Marushina (1978)|
|-Russian Far East||Present||Native||Invasive|
|-Southern Russia||Present, Widespread||Native||Invasive||Original citation: Marushina (1978)|
|-Western Siberia||Present, Widespread||Native||Invasive||Original citation: Marushina (1978)|
|Serbia and Montenegro||Present|
|-Balearic Islands||Present||Native||Invasive||Original citation: Gomez de Aizpurua, 1992|
|Ukraine||Present||Native||Invasive||Original citation: Marushina (1978)|
|United States||Absent, Formerly present||Original citation: Holland (1903)|
|-New York||Absent, Formerly present||Original citation: Holland (1903)|
History of Introduction and SpreadTop of page
Habitat ListTop of page
|Terrestrial||Managed||Cultivated / agricultural land||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Managed||Protected agriculture (e.g. glasshouse production)||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Managed||Managed forests, plantations and orchards||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Managed||Managed grasslands (grazing systems)||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Managed||Disturbed areas||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Managed||Rail / roadsides||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Managed||Urban / peri-urban areas||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Natural forests||Principal habitat||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Natural grasslands||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Riverbanks||Present, no further details||Harmful (pest or invasive)|
|Terrestrial||Natural / Semi-natural||Wetlands||Present, no further details||Harmful (pest or invasive)|
|Littoral||Coastal areas||Present, no further details||Harmful (pest or invasive)|
Hosts/Species AffectedTop of page
Laboratory investigations of nun moth preferences and utilization of Eurasian host plants provide limited and contradictory information (Bejer, 1988). For example, laboratory studies rank the preferred species, spruce, pine and larch, as intermediate to low in food value (Bejer, 1988). Most of the host plant work carried out on L. monacha has concentrated on the relationships between bud burst on the main hosts, P. abies and P. sylvestris, and the hatching of L. monacha larvae. This work has shown that host phenology is as important as host preference in determining the survival and successful development of L. monacha larvae. For example, when the larvae hatch before foliage bud burst, the presence of male flowers or buds on Pinus sp. is critical to larval survival and growth (Bejer, 1988). Keena (2003) tested 26 North American tree species and Withers and Keena (2001) tested Pinus radiata to project the potential host range of this insect if accidentally introduced into North America or New Zealand. The following tree species were found to be suitable hosts for L. monacha: Abies concolor, Picea glauca, Picea pungens, Pinus radiata, Tsuga canadensis, Betula populifolia, Prunus serotina, Quercus lobata, and Quercus velutina.
Host Plants and Other Plants AffectedTop of page
Growth StagesTop of page
SymptomsTop of page
List of Symptoms/SignsTop of page
|Growing point / external feeding|
|Inflorescence / external feeding|
|Inflorescence / webbing|
|Leaves / external feeding|
|Leaves / webbing|
|Stems / external feeding|
|Stems / webbing|
Biology and EcologyTop of page
The haploid karyotype of L. monacha is 31 chromosomes (Seiler and Haniel, 1921). Robinson (1971) provides a review of all the early genetics work on L. monacha, including the inheritance of melanism variation in both the larvae and adults. Recent investigations to characterize L. monacha DNA (Pfeifer et al., 1995; Bogdanowicz et al., 2000) and that of related species provide means of distinguishing L. monacha from closely related species.
Adult L. monacha fly from mid-July to the beginning of September (the exact time depends on the climate of the region). The males are nocturnally active and the females release a pheromone to attract the males. The adults are most active around midnight and the males are much more active than the females. Although the females fly, they usually sit on stems to await the male. Once mated, the females lay from 70 to 300 eggs in clusters of approximately 40 eggs, in bark crevices or under lichens on the bark. After depositing most of her eggs, the female may fly more actively. The L. monacha embryo completes development 2 to 6 weeks after the egg is laid (depending on temperature) and then enters diapause for about 10 weeks. Hatching usually occurs in the beginning of May. First- and second-instars are capable of wind-dispersal over considerable distances. The larvae have five to seven instars and pupation takes place in July. The males typically emerge a few days before the females. This brief summary was prepared using summaries by Bejer (1988), Grijpma (1989), and Lipa and Glowacka (1995).
L. monacha is a typical transpalearctic species with a wide distribution from Japan, Korea, China, throughout Russia (southern parts of Russia Far East, Eastern and Western Siberia, Southern Urals, European part; see map in Baranchikov, 1997), and most European countries. It occurs within a band between northern latitudes 43°N and 57°N (Carter, 1984) including southern England, Denmark, Sweden and Finland in the north, and Spain, Portugal and Italy at elevations of 1000 to 2000 m in the south (Lipa and Glowacka, 1995). The zone in which periodic outbreaks occur is generally bounded by the July isotherm of 16°C and September isotherm of 10.5°C (Bejer, 1988). The outbreak areas are semiarid and several studies have indicated that more frequent outbreaks occur in drought-sensitive sites where the hosts are probably stressed (Bejer, 1988; Lipa and Glowacka, 1995).
Natural enemiesTop of page
|Natural enemy||Type||Life stages||Specificity||References||Biological control in||Biological control on|
|Agria housei||Parasite||Arthropods|Larvae; Arthropods|Pupae|
|Bacillus thuringiensis galleriae||Pathogen||Arthropods|Larvae|
|Bacillus thuringiensis kurstaki||Pathogen||Arthropods|Larvae|
|Bacillus thuringiensis subsp. dendrolimus||Pathogen||Arthropods|Larvae|
|Bacillus thuringiensis thuringiensis||Pathogen||Arthropods|Larvae|
|Calosoma sycophanta||Predator||Arthropods|Larvae; Arthropods|Pupae|
|Compsilura concinnata||Parasite||Arthropods|Larvae; Arthropods|Pupae|
|Cytoplasmic polyhedrosis virus (CPV)||Pathogen||Arthropods|Larvae|
|cytoplasmic polyhedrosis viruses||Pathogen||Arthropods|Larvae|
|Eurytoma verticillata||Parasite||Arthropods|Larvae; Arthropods|Pupae|
Notes on Natural EnemiesTop of page
Several species of birds, arthropods and small mammals are reported to prey on the eggs, larvae, pupae and adults of L. monacha (Steinfatt, 1942). Birds are considered to be the main predators of both eggs and larvae, but may not have much influence during outbreaks (von Wellenstein and Schwenke, 1978).
Means of Movement and DispersalTop of page
Newly hatched L. monacha larvae often climb to the top of trees and can become wind blown, aided by the silk threads they produce and the specialised hairs they possess (see Morphology section for description). The larvae may also crawl to new hosts when their immediate food supply has been exhausted. Both the male and female moths fly, although females usually fly less until they have laid most of their eggs, thus contributing to the expansion of outbreaks over sequential years (Bejer, 1988).
Movement in Trade
The adults are readily attracted to lights (Wallner et al., 1995) and have been observed in the vicinity of ports where they could lay eggs in or on structures that will be transported. Whole bolts of preferred host with intact bark could also contain hidden eggs. These eggs would not be readily spotted because the females lay their eggs under bark scales and in cracks. In outbreak areas, other stages could potentially become associated with items that will be moved in trade or vehicles.
Plant TradeTop of page
|Plant parts liable to carry the pest in trade/transport||Pest stages||Borne internally||Borne externally||Visibility of pest or symptoms|
|Bark||eggs||Yes||Yes||Pest or symptoms usually visible to the naked eye|
|Flowers/Inflorescences/Cones/Calyx||larvae||Yes||Pest or symptoms usually visible to the naked eye|
|Leaves||larvae||Yes||Pest or symptoms usually visible to the naked eye|
|Stems (above ground)/Shoots/Trunks/Branches||adults; eggs; larvae; pupae||Yes||Pest or symptoms usually visible to the naked eye|
Impact SummaryTop of page
|Fisheries / aquaculture||None|
|Fisheries / aquaculture||None|
ImpactTop of page
Environmental ImpactTop of page
Social ImpactTop of page
DiagnosisTop of page
Detection and InspectionTop of page
Similarities to Other Species/ConditionsTop of page
L. monacha owes its common name (nun moth) to its similarity to another moth, the monk (Panthea coenobita), which also has a black and white monk-cloak colouring (Bejer, 1988). P. coenobita (Noctuidae) differs from L. monacha (Lymantriidae) in that it has ocelli and tufts of scales on the dorsum of the thorax. They also belong to different Lepidoptera families.
Lymantria minomonis okinawaensis found on Okinawa and Lymantria minomonis sugii in Taiwan have previously been misidentified as L. monacha because they look like the lighter form (Kishida, 1987).
In the Russian Far East, adult male L. monacha held with forceps with their wings positioned above the body can produce a clear sound similar to one of Lymantria mathura (Baranchikov et al., 2004). This has never been reported from Siberian and European populations of L. monacha.
Prevention and ControlTop of page
Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.
The tendency of forestry to move toward large areas of monoculture and even-aged stands tends to increase the incidence of L. monacha outbreaks. Another factor is the planting of trees in areas not well suited to the tree species, thus stressing the trees and increasing their susceptibility to pest attack. To reduce the risk of large-scale outbreaks, Bejer (1988) suggests that spruce forests in the outbreak zone or on susceptible sites should be well dispersed, removed, or put on a shorter rotation. Establishing mixed stands of conifers or adding deciduous trees might also reduce the risks of outbreak (Bejer, 1988). Neither of these options would be easy to implement for economic reasons.
At present, only Bacillus thuringiensis products are available for operational use against L. monacha. There is some variation in the control results obtained for different formulations (Glowacka, 1989; Glowacka, 1995). B. thuringiensis has been widely used with good success in Germany (Altenkirch et al., 1986; Langenbruch, 1993), Russia (Bakhvalov et al., 1984; Marchenko, 1995), Belarus (Krushev and Marchenko, 1981) and the Czech Republic (Svestka, 1995).
For more than a century it has been known that natural epizootics of the nuclear polyhedrosis virus are the main factor that causes the collapse of L. monacha outbreaks (Grijpma, 1989). Several attempts to produce it have been made but no large-scale production for operational use has yet resulted.
Mating disruption using micro encapsulated pheromones sprayed on trees has been attempted and shown to reduce population numbers even in the following year (Vrkoch et al., 1981; Jensen, 1983).
Chemical controls have been used against L. monacha since 1892, when the world's first synthetic insecticide was applied (Ferguson, 1992). The large-scale use of a number of chlorinated hydrocarbons followed until they were banned. Synthetic pyrethroids and growth inhibitors are still in use against L. monacha. The growth inhibitors are slow to act so can only be used when the populations are low enough or caught early enough so that the growing tips of the conifers are not completely destroyed. Synthetic pyrethroids have negative effects on non-target organisms so they cannot be used in environmentally sensitive areas, but they are fast-acting and cheap, so they are still used.
Field Monitoring/Economic Threshold Levels
Bogenschütz (1982) proposed a three-stage monitoring system for determining when control treatments are needed: use pheromone warning traps to survey endemic populations; a more intense survey when the numbers of males trapped reaches a critical threshold; and forecasting larval damage to determine if treatments are needed. Schmutzenhofer (1986) suggested that 2000 to 3000 males per trap over the entire flight season is the threshold warning of an outbreak in the next year. Skatulla (1989) demonstrated a correlation between pheromone moth catches and population density and proposed a critical threshold value of 60 to 70 males per trap per night. Markov (1999) gave population density thresholds for control actions against L. monacha in Russia.
Jensen (1991) proposed a tentative integrated pest management programme for L. monacha. More work on thresholds for intervention using different methods still needs to be done to make this possible.
ReferencesTop of page
Anonymous, 1991. Pest risk assessment on the importation of larch from Siberia and the Soviet Far East. USDA Forest Service, Miscellaneous Publication No. 1495, 280 pp.
Baranchikov YuN, 1997. Siberian forest insects: ready for export. Exotic Pests of Eastern Forests. Conference Proceedings. Nashville: USDA Forest Service, 85-91. http://www.invasive.org/symposium/baranchi.html.
Baranchikov YuN, Wallner WE, Carde RL, Turova GI, Hamble LM, Yurchenko GI, 2004. Dynamics of Lymantriid moths flight to the electrical lights in the Primorskiy Kray - two years observations. Entomologicheskiye Issledovaniya v Sibiri. Vol. 3. Krasnoyarsk: Institut lesa SO RAN, 84-99.
Bejer B, 1988. The nun moth in European spruce forests. In Berryman AA, ed. Dynamics of forest insect populations: Patterns, causes, implications. New York, USA: Plenum Press, 211-231.
Bryk F, 1934. Lymnatriidae. In: Strand, ed. Lepidopterorum Catalogus, 62:1-441. Berlin, Germany: W Junk.
Carter DJ, 1984. Pest Lepidoptera of Europe, with special reference to the British Isles. Series Entomologica, Volume 31.
Chao C-L, 1978. Memoirs of Chinese Economic Entomology, Volume 12. Lepidoptera, Lymantriidae. Peking, China: Scientific Publication Center, 1-121.
Dyar H, 1897. A generic revision of the Hypogymnidae (Liparidae). Canadian Entomologist, 29:12-16.
Dzutevski B, Cakar L, 1955. The mass appearance of L. monacha in the Beech forests of Macedonia. [Masovna pojava Lymantria monacha L. u. bukovim sumama Makedonije.] Zast. Bilja No. 32, (55-60). 2 refs.
EPPO, 2014. PQR database. Paris, France: European and Mediterranean Plant Protection Organization. http://www.eppo.int/DATABASES/pqr/pqr.htm
Ferguson DC, 1978. The moths of America north of Mexico. Fascicle 22.2 Noctuoidea, Lymantriidae. The moths of America north of Mexico. Fascicle 22.2 Noctuoidea, Lymantriidae. E.W. Classey Ltd and the Wedge Entomological Research Foundation. London UK, 110 + x pp.
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Glowacka B, 1989. Pathogenic viruses and bacteria of the nun moth (Lymantria monacha L.) during the outbreak 1978-1984 in Poland. In: Wallner WE, McManus KA, Technical Coordinators. Proceedings, Lymantriidae: A comparison of features of New and Old World tussock moths. USDA Forest Service, Northeastern Forest Experiment Station, General Technical Report NE 123, 401-415.
Glowacka B, 1995. Mortality of nun-moth (Lymantria monacha L.) and non-target arthropods in control treatments of the 1994 campaign. Folia Forestalia Polonica. Seria A, LeSnictwo, No. 37:89-109; 9 ref.
Gries G, Gries R, Khaskin G, Slessor KN, Grant GG, Liska J, Kapitola P, 1996. Specificity of nun and gypsy moth sexual communication through multiple-component pheromone blends. Naturwissenschaften, 83(8):382-385; 20 ref.
Grijpma P, 1989. Overview of research on Lymantrids in eastern and western Europe. In: Wallner WE, McManus KA, Technical Coordinators. Proceedings, Lymantriidae: A comparison of features of New and Old World tussock moths. USDA Forest Service, Northeastern Forest Experiment Station, General Technical Report NE 123, 21-49.
Hampson GF, 1892. The Fauna of British India, including Ceylon and Burma. Moths. Volume 1. London, UK: Taylor and Francis, 527 pp.
Herting B, Simmonds FJ, 1976. A catalogue of parasites and predators of terrestrial arthropods. Section A. Host or prey/enemy. Volume VII. Lepidoptera, Part 2 (Macrolepidoptera). A catalogue of parasites and predators of terrestrial arthropods. Section A. Host or prey/enemy. Volume VII. Lepidoptera, Part 2 (Macrolepidoptera). Commonwealth Agricultural Bureaux. Farnham Royal UK, [5 +]221 pp.
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Inoue H, 1957. A revision of the Japanese Lymantriidae (II). Japanese Journal of Medical Science and Biology 10:187-219.
Keena MK, Shields KS, Torsello M, 1998. Nun moth: potential new pest. USDA Forest Service, Northeastern Area NA PR 95 98. Radnor, PA.
Kim CW, Nam SH, Lee SM, 1982. Illustrated Flora and Fauna of Korea Vol. 26, Insecta 8. Seoul, Korea Republic: Ministry of Education.
Kirby W, 1892. A Synonymic Catalogue of Lepidoptera Heterocera (Moths). London, UK: Gurney & Jackson.
Kishida Y, 1987. A new subspecies of Lymantria minomonis Matsumura (Lymantriidae) from Okinawa I. Japan Heterocerists’ Journal, 144:292-293.
Klimetzek D, 1979. Pine insects in south Germany: a historical entomological study. [Kieferninsekten in Suddeutschland. Eine historisch-entomologische Studie.] Forstwissenschaftliches Centralblatt, 98(5):277-280; 6 ref.
Kolomiets NG, 1990. Insects, predators and parasites of the nun moth (Lymantria monacha L., Lepidoptera) in the USSR. Fauna Sibiri. Nasekomyye i gel’minty. Novosibirsk: Nauka Publication, 242-251.
Kolubajiv O, 1962. Ergebnisse der Zuchten von Entomophagen (der Parasiten und Räuber) der Schädlichen Insekten (vorwiegent der Forstschädlinge) der Zeitperiode von 1934 bis 1958. Rozpravy Ceskoslovenske Akademie Ved, 72:1-72.
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